Nutrients cause grassland biomass to outpace herbivory

E. T. Borer, W. S. Harpole, P. B. Adler, C. A. Arnillas, M. N. Bugalho, M. W. Cadotte, M. C. Caldeira, S. Campana, C. R. Dickman, T. L. Dickson, I. Donohue, A. Eskelinen, J. L. Firn, P. Graff, D. S. Gruner, R. W. Heckman, A. M. Koltz, K. J. Komatsu, L. S. Lannes, A. S. MacDougallJ. P. Martina, J. L. Moore, B. Mortensen, R. Ochoa-Hueso, H. Olde Venterink, S. A. Power, J. N. Price, A. C. Risch, M. Sankaran, M. Schütz, J. Sitters, C. J. Stevens, R. Virtanen, P. A. Wilfahrt, E. W. Seabloom

Research output: Contribution to journalArticlepeer-review

37 Scopus citations


Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs (‘consumer-controlled’). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food (‘resource-controlled’). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk.

Original languageEnglish (US)
Article number6036
JournalNature communications
Issue number1
StatePublished - Nov 27 2020

Bibliographical note

Funding Information:
This work was generated using data from the Nutrient Network ( experiment, funded at the site scale by individual researchers. Author contributions are detailed in the “Author contributions” section and Supplementary Table 5; Supplementary Table 6 lists all data contributors who are not authors. Coordination and data management have been supported by funding to E.T.B and E.W.S. from the National Science Foundation Research Coordination Network (NSF-DEB-1042132) and Long-Term Ecological Research (NSF-DEB-1234162 and NSF-DEB-1831944 to Cedar Creek LTER) programs, and the Institute on the Environment (DG-0001-13). We also thank the Minnesota Supercomputer Institute for hosting project data and the Institute on the Environment for hosting Network meetings. The authors dedicate this paper to the memory of Enrique Chaneton who contributed data and early ideas to this work.

Publisher Copyright:
© 2020, The Author(s).

Copyright 2021 Elsevier B.V., All rights reserved.


  • Biomass
  • Confidence Intervals
  • Fertilizers
  • Grassland
  • Herbivory/physiology
  • Nitrogen/analysis
  • Phosphorus/analysis
  • Time Factors

PubMed: MeSH publication types

  • Research Support, Non-U.S. Gov't
  • Research Support, U.S. Gov't, Non-P.H.S.
  • Journal Article


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